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RESEARCH ARTICLE
How do land-use legacies affect ecosystem services in UnitedStates cultural landscapes?
Carly Ziter . Rose A. Graves . Monica G. Turner
Received: 30 October 2016 /Accepted: 6 June 2017
� Springer Science+Business Media B.V. 2017
Abstract
Context Landscape-scale studies of ecosystem ser-
vices (ES) have increased, but few consider land-use
history. Historical land use may be especially impor-
tant in cultural landscapes, producing legacies that
influence ecosystem structure, function, and biota that
in turn affect ES supply.
Objectives Our goal was to generate a conceptual
framework for understanding when land-use legacies
matter for ES supply in well-studied agricultural,
urban, and exurban US landscapes.
Methods We synthesized illustrative examples from
published literature in which landscape legacies were
demonstrated or are likely to influence ES.
Results We suggest three related conditions in which
land-use legacies are important for understanding
current ES supply. (1) Intrinsically slow ecological
processes govern ES supply, illustrated for soil-based
and hydrologic services impaired by slowly processed
pollutants. (2) Time lags between land-use change and
ecosystem responses delay effects on ES supply,
illustrated for biodiversity-based services that may
experience an ES debt. (3) Threshold relationships
exist, such that changes in ES are difficult to reverse,
and legacy lock-in disconnects contemporary land-
scapes from ES supply, illustrated by hydrologic
services. Mismatches between contemporary land-
scape patterns and mechanisms underpinning ES
supply yield unexpected patterns of ES.
Conclusions Today’s land-use decisions will gener-
ate tomorrow’s legacies, and ES will be affected if
processes underpinning ES are affected by land-use
legacies. Research priorities include understanding
effects of urban abandonment, new contaminants, and
interactions of land-use legacies and climate change.
Improved understanding of historical effects will
improve management of contemporary ES, and aid
in decision-making as new challenges to sustaining
cultural landscapes arise.
Keywords Land-use change � Urban ecosystems �Exurban ecosystems � Agricultural ecosystems �Historical ecology
Introduction
Landscape-scale studies of ecosystem services (ES)
have increased in recent years (Maes et al. 2012;
Mitchell et al. 2015a), including in the US (Nelson
et al. 2009; Qiu and Turner 2013; Carpenter et al.
2015a; Blumstein and Thompson 2015), but relatively
little attention has been paid to the role of land-use
history. Many studies map ES supply (and ES
tradeoffs and synergies) across the landscape (Bur-
khard et al. 2013; Malinga et al. 2015) as an important
C. Ziter (&) � R. A. Graves � M. G. Turner
Department of Zoology, University of Wisconsin-
Madison, Madison, WI 53717, USA
e-mail: ziter@wisc.edu
123
Landscape Ecol
DOI 10.1007/s10980-017-0545-4
first step towards incorporating ES into policy (Chan
et al. 2006; Naidoo et al. 2008; Raudsepp-Hearne et al.
2010). However, these maps often rely on current
land-cover pattern, which is a poor surrogate for ES
supply (Eigenbrod et al. 2010; Schulp et al. 2014), but
underlying processes are often not well known.
Understanding the mechanisms driving landscape
patterns of ES is increasingly important as ES become
explicitly incorporated in decision-making; for exam-
ple, the Obama administration issued a 2015 memo-
randum directing all US federal agencies to
incorporate ES into their planning (Donovan et al.
2015). Spatial and temporal drivers of ES supply must
be considered to improve mechanistic understanding
of ES supply at landscape scales, particularly in
human-dominated landscapes. However, most analy-
ses to date have estimated ES supply without
addressing the role of land-use legacies—the persis-
tent effects of past events, patterns, or conditions on
the contemporary landscape.
Historical land use produces legacies that influence
the structure, function, and biota of many ecosystems
(Foster et al. 2003; Lunt and Spooner 2005; Burgi et al.
2017), which in turn affect ES supply. Thus, both
contemporary and historical land-use patterns may
influence ES supply. In the Southern Appalachian
Mountains for example, where land use change is
shaped by transition to a nature-based economy,
effects of land-use history have been well documented
on soil nutrients (Fraterrigo et al. 2005) and the
presence and abundance of native (Elliott et al. 2014)
and non-native forest understory plant species (Kuh-
man et al. 2011, 2013). These land-use legacies in soils
and vegetation in turn influence opportunities for
wildflower viewing, a cultural ecosystem service that
attracts tourists, and for which the region is widely
known (Graves et al. 2017). In Sheffield, UK,
historical land-use patterns were of greater importance
than current patterns in explaining variability in
several indicators of contemporary ES supply, includ-
ing aboveground carbon storage, recreational use, and
bird species richness (Dallimer et al. 2015). Recent
studies also suggest effects of historical land use on ES
may change over time (Watson et al. 2014). ES
dynamics can differ markedly within the same land-
scape following recovery from disturbance (Suther-
land et al. 2016), and relationships among sets of ES
can vary qualitatively over time with land-use history
(Renard et al. 2015; Tomscha and Gergel 2016).
Understanding when land-use legacies explain
variation in contemporary ES is particularly relevant
in cultural landscapes—landscapes that are shaped by
human activities and possess features valued by
humans (Schaich et al. 2010; Plieninger et al. 2014).
Cultural landscapes have long histories of land-
use/land-cover (LULC) change as a result of their
deliberate management, and are also areas of high ES
demand. While studies have demonstrated lasting
effects of land-use legacies in cultural landscapes
globally (e.g., Cousins and Eriksson 2002; Burgi and
Gimmi 2007; Plieninger et al. 2010; Dullinger et al.
2013; Dallimer et al. 2015), we focus specifically on
land-use legacies and ES in cultural landscapes of the
United States. We consider contemporary agricultural,
urban and exurban landscapes (Fig. 1) that represent
areas most people live and work (Brown et al. 2005;
Theobald 2005; Radeloff et al. 2012; Groffman et al.
2014), though we recognize land-use legacies also
affect the ecology of other US landscapes (Foster et al.
2003). Sometimes referred to as ‘‘vernacular cultural
landscapes’’, these landscapes have evolved uninten-
tionally as a result of everyday human use over time—
thus reflecting the physical, biological, and cultural
character of everyday lives (Alanen and Melnick
2000; The Cultural Landscape Foundation). Cultural
landscapes can also encompass heritage areas shaped
by particular religious, racial, or cultural groups
(‘‘ethnographic landscapes’’), consciously designed
landscapes such as parks or campuses (‘‘designed
landscapes’’), and landscapes associated with historic
events (‘‘historic sites’’) (The Cultural Landscape
Foundation, National Park Service); however the
types of cultural landscapes we consider here currently
make up the majority of cultural landscapes in
America (Alanen and Melnick 2000).
Prior to European settlement US landscapes were
largely forested, but also contained significant prairie
and savanna cover—with landscape and vegetation
patterns driven in part by indigenous peoples, partic-
ularly through centuries of fire management (Macleish
1994; Delcourt and Delcourt 2004). Land use changed
profoundly in the US following European settlement,
although the timing and sequence of land-use changes
varied among regions (Whitney 1994; Turner et al.
1998). The seventeenth through nineteenth centuries
saw widespread forest loss, primarily as a conse-
quence of farming, the burgeoning lumber industry,
and widespread fuelwood collection. Occurring in
Landscape Ecol
123
many regions at a pace much faster than analogous
forest clearance in Europe, this conversion to culti-
vated land resulted in fragmentation of North Amer-
ican woodlands (Whitney 1994). Cropland peaked in
the US in the 1940s and has since fluctuated around
162million ha (Turner et al. 1998). Some areas cleared
for agriculture have reforested in the past century due
to lack of cultivation (e.g., New England, Southeast,
and upper Midwest), whereas clearing for agriculture
was more permanent in other regions (e.g., lower
Midwest) (Whitney 1994). In contemporary cultural
landscapes of the US, agricultural areas remain
dominated by croplands and pastures (Fig. 1a) while
urban landscapes are characterized by high-density
commercial, industrial, and residential land use
(Fig. 1b). Exurban landscapes are composed of low-
density residential development, often in rural areas
that offer environmental amenities or on former
croplands surrounding mid-sized cities (Fig. 1b).
Exurban land use occupies more area than cities in
the US and is increasing rapidly (Theobald 2005;
Brown et al. 2005). The relatively recent historical
context (*150 years) sets these US landscapes apart
from the iconic cultural landscapes of Europe, where
land-use patterns have been more stable over much
longer periods of time (Keatley et al. 2011).
Cultural landscapes are closely linked to human
wellbeing through the services they provide, and like
such landscapes elsewhere around the world, they are
dynamic (Plieninger et al. 2014). Subject to shifts in
land cover and often changing in response to policy or
economics, these anthropogenically modified land-
scapes are defined by transition from a previous LULC
type to their current state. Thus, ES provided by these
landscapes may be particularly influenced by histor-
ical land-use patterns. As landscapes continue to
transition towards greater use and dominance by
people (Foley et al. 2005; Ellis 2011), understanding
how land-use history influences ES supply and how
current activities may produce future land-use legacies
is necessary for managing cultural landscapes
sustainably.
Although some recent research examines effects of
historical land use on ES supply, studies that incor-
porate historical information are rare in the ecosystem
services literature. It is often difficult to locate
consistent data sources, some of which are qualitative
and most of which have not been assembled over
whole landscapes. New methods for incorporating
historical aspects of ES are emerging (Tomscha et al.
2016), yet the difficulty and cost of assembling the
data sources make it unrealistic for every study to
consider land-use legacies for each ES in question.
Knowing when it is most important to account for
history in estimating ES supply is an important step
towards understanding patterns of ES provision.
Here, we present a framework for considering when
land-use legacies matter for ES supply in US cultural
landscapes, drawing on illustrative examples from
well-studied agricultural, urban, and exurban land-
scapes. We suggest three related but distinct condi-
tions in which land-use legacies are especially
Fig. 1 Photos illustrating different types of cultural landscape
in the United States. a An agricultural landscape in the Yahara
watershed, southern Wisconsin. Photo credit: University of
Wisconsin-Madison Water Sustainability and Climate project.
b An urban landscape in Madison, Wisconsin. Photo credit:
University of Wisconsin-Madison water sustainability and
climate project. c An exurban landscape in the southern
Appalachian Mountains of western North Carolina. Photo
credit: US Long-term Ecological Research Network Office
Landscape Ecol
123
important for understanding current ES supply: (1)
intrinsically slow ecological processes govern ES
supply; (2) time lags between land-use change and
ecosystem responses delay the effects on ES supply;
and (3) threshold relationships exist, such that changes
in ES may be difficult to reverse. These three
conditions are not mutually exclusive, but their
influence on ES can be distinct as highlighted by
examples presented herein. In all cases, the mismatch
between what is suggested by the contemporary
landscape and mechanisms underpinning ES supply
can yield unexpected patterns of ES. We conclude by
considering the influence that current land use may
have on future ES supply.
Land-use legacies and contemporary ES supply
Slow processes govern ES supply
Land-use legacies are important for ES when services
are underlain by intrinsically slow ecological pro-
cesses (Fig. 2a) (Walker et al. 2012). Soil-based ES
offer useful examples because many ecological
processes and soil nutrient stocks are slow to recover
from past land use. For example, soil carbon
(C) stocks are often used as an indicator of climate
regulation services, as well as water retention and
general soil fertility, and soil C stocks in urban and
exurban landscapes vary with historical land-use and
time since development (Golubiewski 2006; Lewis
et al. 2006; Raciti et al. 2011). Interestingly, effects
of land-use legacies on soil C in cities vary in
magnitude and direction among regions. In Balti-
more, Maryland (mesic mid-Atlantic coast), residen-
tial lots developed on former agricultural land had
initially low C stocks compared to those converted
from forest, because years of agricultural use
depleted soil C (Raciti et al. 2011). Following
development, former agricultural lands accumulated
soil C over time, whereas former forests did not
(Raciti et al. 2011). Contrastingly, in the greater
Phoenix area, Arizona (xeric Southwest), residential
lots developed on former agricultural lands had
substantially higher soil C stocks compared to those
converted from native desert. In arid regions, soil C
was increased during years of agricultural use, rather
than depleted, in response to irrigation and nutrient
augmentation (Lewis et al. 2006). Differences in soil
C based on land-use history persisted for decades
after residential development because processes that
change soil C pools are inherently slow. Although
effects of land-use history differed in important ways
between regions, variation in present-day soil C
(a)
(b)
(c)
Fig. 2 Three conditions in which land-use legacies are
important for understanding current ecosystem service (ES)
supply. a Intrinsically slow ecological processes govern ES
supply. b Time lags between land-use change and ecosystem
responses delay effects on ES supply. c Threshold relationshipsexist, such that changes in ES are difficult to reverse, and legacy
lock-in disconnects contemporary landscapes from ES supply.
For each of the three conditions, dashed lines represent possible
trajectories of ES supply following land-use change
Landscape Ecol
123
stocks could not be explained in the absence of
understanding land-use history.
Water quality and associated hydrologic services
(Brauman et al. 2007) can be similarly influenced by
land-use legacies associated with long-term persis-
tence of nutrients or environmental pollutants long
after inputs have ceased. In this case, the rate
ecosystems can process or purge a pollutant is
intrinsically slow. Consider soil phosphorus (P), a
common pollutant associated with eutrophication of
inland lakes and widely used indicator of lake water
quality and hydrologic services (Carpenter et al. 1998;
Lathrop 2007). Fertilizer and manure application
increase soil P, often over decades of farming.
Because P adheres to soil particles and is not readily
soluble, it is transported very slowly from croplands to
surface waters via soil erosion and runoff, much of
which occurs during high-intensity rain events (Car-
penter et al. 2015b). Drawdown of soil P is very slow
in agricultural landscapes (Hamilton 2011), and over-
fertilized soils remain P enriched for decades (Bennett
et al. 1999) even following implementation of con-
servation strategies and/or land-use change aimed at P
reduction (Hamilton 2011; Sharpley et al. 2013). In the
YaharaWatershed of southernWisconsin, a century of
agricultural P inputs have resulted in spatially variable
but highly elevated soil P levels that continue to act as
a hidden driver of water quality (Bennett et al. 2005;
Kara et al. 2011). Persistent efforts to control P
pollution have produced negligible improvements to
water quality (Gillon et al. 2016), and expanding
urbanization on former croplands continues to release
soil P in urban runoff (Betz et al. 2005). Land-use
history is a key driver of soil P that is not necessarily
apparent or explained by contemporary LULC
patterns.
Environmental pollutants such as mercury and
organochlorine pesticides (e.g., DDT, PCBs) also
persist in cultural landscapes for decades to centuries
as a legacy of industrial, agricultural, and residential
use (Wang et al. 2004; Weber et al. 2008). Contam-
inated water and sediments enter the food web at low
trophic levels and bioaccumulate in higher trophic
organisms such as fish; the resultant reduction in
seafood quality impacts provisioning and cultural ES
(Holmlund and Hammer 1999). Loss of these pollu-
tants from water and sediments through degradation or
volatilization is a slow process. Consider the densely
populated San Francisco Bay watershed in California.
Although regional organochlorine pesticide use
peaked 30–40 years ago, pesticide residues remain
high enough to warrant current sport fish consumption
advisories (Connor et al. 2007; Davis et al. 2007;
Schoellhamer et al. 2007). Comparable trends are seen
in urban and ex-urban landscapes encompassing the
Houston Ship Channel system in Texas, where con-
taminated sediments from historical dioxin use are the
primary contributor to seafood consumption advi-
sories for several species (Dean et al. 2009), and in the
city of Oak Ridge, Tennessee, where legacy mercury
continues to accumulate in freshwater fish as a result
of streambank erosion (Southworth et al. 2011).
Similarly to P, these legacy pollutants are likely to
endure for decades despite substantial reduction in
inputs. Model predictions suggest a 10–30 year period
for pesticide loads to reach risk-reduction goals under
scenarios of zero pesticide loading in San Francisco
Bay (Connor et al. 2007), while in Oak Ridge, fish
have sustained high concentrations of mercury and
PCBs for decades despite discontinued use and
multiple remediation efforts (Southworth et al.
2011). Thus, when processes that mitigate land-use
legacies associated with an ES are intrinsically slow,
land-use history will affect ES supply.
Time lags delay effects of land use on ES supply
Time lags between land-use change and ecological
responses can cause delayed effects on ES that are
better explained by historical land use than current
landscapes (Raudsepp-Hearne et al. 2010). The ulti-
mate change in ES may occur slowly, similarly to the
previous examples, or relatively quickly, but a key
factor is a notable delay between land-use change and
the ES response (‘‘lagged response’’ in Fig. 2b). Time
lags occur when, for a time period following land-use
change, ES supply remains consistent with supply
prior to land-use change, or when the rate of change in
ES supply increases or decreases over time following
land-use change, even in absence of further land-use
change. Such delayed effects may be especially
pronounced for biodiversity-based services (Kremen
2005) because historical habitat loss or fragmentation
caused by land-use changes can lead to extinction
debts wherein species loss continues beyond the initial
event or driver (Tilman et al. 1994; Jackson and Sax
2009; Essl et al. 2015). Extinction debts can be
assumed if historical land use, habitat area and
Landscape Ecol
123
connectivity better explain current species distribution
than do current land use and landscape variables. From
an ES perspective, an extinction debt can substantially
delay the effects of land-use change on supply of
biodiversity-based ES, such as C storage, pollination,
pest control, and nature viewing. Dependent upon the
resultant change in species composition and the
particular ES, these effects may be negative or
beneficial, and may occur as a result of changes in
both native and invasive species. Consequences may
be particularly large if species affected act as ecosys-
tem service providers (Luck et al. 2009).
Time lags between human LULC change and
species loss may help explain differences in ES supply
that are unexplained by current LULC patterns
(Fig. 2b). Ignoring potential extinction debts or col-
onization patterns may lead to overly optimistic
assessments of such ES, or underestimation of ES
supply, if species changes influence ES positively.
Cultural landscapes in which natural and semi-natural
habitats are reduced or fragmented may be especially
susceptible to future species extinctions and perma-
nent loss of certain biodiversity-based ES (i.e.,
biodiversity-based ES debt, Isbell et al. 2014). Despite
the potential for biodiversity and ecosystem function
lags in response to land-use change (Valiente-Banuet
et al. 2014), time lags have been largely absent from
ES research (but see Dallimer et al. 2015).
Perennial plant communities often display time lags
between habitat loss or degradation and eventual
species loss (Helm et al. 2005) which can lead to
biodiversity-based ES extinction debts. In the agri-
cultural and exurban landscapes of the Midwestern
US, prairie remnants provide multiple services. Con-
sistent with predictions of extinction debt, extinction
rates in prairie remnants have accelerated over time,
suggesting a lagged response of community diversity
to land-use changes; present day extinction and
colonization in prairie remnants is strongly related to
historical land use (Alstad et al. 2016). Historical land
management in prairie remnants explained coloniza-
tion by non-native species and extinction of native
species; loss of native species can impact ES in
surrounding agricultural lands by affecting the diver-
sity and identity of beneficial arthropods such as
predators, parasitoids, and pollinators. Indeed, exper-
imental work identified distinct assemblages of ben-
eficial arthropods associated with different native
prairie plant species (Bennett and Gratton 2013). In
addition to extinction debts for ES such as pollination
and pest control, lagged species loss or gains in prairie
and grassland habitats can lead to unpredicted change
in above-ground and below-ground C stocks (Isbell
et al. 2011). In the Midwestern US prairie remnants,
historical land-use patterns explained shifts from
perennial species to annual species in the prairie
remnants (Alstad et al. 2016) and eventual reductions
in C storage.
Time lags in biodiversity-based ES can exist due to
slow processes of re-colonization that depend on
historical land use and connectivity (Lindborg and
Eriksson 2004; Gonzalez et al. 2009; Aguirre-
Gutierrez et al. 2015). Current land-use patterns and
connectivity strongly affect supply and delivery of ES
(Kremen et al. 2007; Cardinale et al. 2012; Mitchell
et al. 2013, 2015b), but some effects of habitat
fragmentation on ES (e.g., C and N retention,
productivity, pollination) can be delayed up to a
decade (Haddad et al. 2015). Effects of historical
habitat fragmentation can persist even after connec-
tivity is re-established. In formerly fragmented long-
leaf pine (Pinus palustris) forests on the southeastern
US coastal plain, current understory plant community
composition on previously agricultural sites was
explained by historical connectivity (Brudvig and
Damschen 2010). In addition, historical land use was
an important factor in predicting the presence and
abundance of the highly invasive fire ant (Solenopsis
invicta) (Stuhler and Orrock 2016), which can cause
acute health issues in human populations as well as
major economic and ecological impacts (Kemp et al.
2000).
Similarly, in amenity-based landscapes of the
eastern US, time lags in understory herb recovery
following agricultural abandonment and eventual
forest regeneration can produce lagged responses in
associated ES (e.g., nature viewing, wildflower
photographing, wild edible or medicinal plants;
Graves et al. 2017). In the southern Appalachian
Mountains, where ecotourism is economically
important, lower richness and abundance of charis-
matic wildflower species in post-agricultural forests
is only partially explained by current forest patch
size, suggesting time lags in re-colonization (Pearson
et al. 1998; Mitchell et al. 2002). In exurban
landscapes in New York, species richness of under-
story herbs in post-agricultural forests depended on
historical isolation and the effect persisted up to
Landscape Ecol
123
100 years after agricultural abandonment (Flinn and
Marks 2004).
Time lags between human-driven land-use change,
ecosystem function and resulting ES are not unique to
biodiversity-based ecosystem services. Hydrologic
services may be particularly affected by time lags in
cultural landscapes, in many cases linked to the slow
processes discussed above. For example, time lags can
range from months to years to decades before changes
in landscape pattern improve water quality in non-
point source watersheds (Meals et al. 2010). Ground-
water in agricultural landscapes can have high pollu-
tant concentrations, and groundwater travel time
coupled with biogeochemical processes leads to time
lags of decades between changes in agricultural
landscapes and corresponding changes in water qual-
ity (Meals et al. 2010; Hamilton 2011). In Iowa,
despite changes in agricultural practices between the
1970s and 2000, groundwater nitrate–N concentra-
tions measured in the early 2000s remained strongly
influenced by 1970s land use (Tomer and Burkart
2003). Actual lag times vary by ES and mechanism,
but ES models based solely on current LULC patterns
will be inaccurate if ES demonstrated lagged
responses and historical land use is not considered.
Surpassing a threshold leads to legacy lock-in
Land-use legacies will be important for understanding
ES supply when threshold relationships exist, and
historical land use produced changes in ES supply that
are difficult to reverse (Fig. 2c). Crossing thresholds
that cannot readily be reversed will effectively
disconnect ES supply from the contemporary land-
scape. However, the potential for threshold dynamics
to influence ES supply in cultural landscapes has
received relatively little consideration, and to our
knowledge, studies have not yet addressed thresholds
and reversibility related to land-use legacies. A variety
of ecological variables exhibit threshold responses as
levels of driver variables change. For example, the
likelihood of a species being present in a habitat patch
can show a threshold response to patch size (Pereira
et al. 2004), and the likelihood of habitat being
connected across a landscape increases rapidly once a
threshold of habitat abundance is passed (Andren
1994). Supply of some ES may show threshold
responses to land-cover abundance in agricultural
landscapes, e.g., indicators of water quality vs. percent
cropland in sub-watersheds of the Yahara Watershed,
Wisconsin (Qiu and Turner 2015) or lake-water clarity
and percent of cropland in riparian zones ofWisconsin
lakes (Rose et al. 2017). For other services, thresholds
associated with socially desirable ES supply, such as
health standards for concentrations of pollutants, may
be surpassed. Some thresholds are reversible; in
others, hysteresis can lead to alternative states that
are very difficult to reverse. In such cases, slow
processes (Fig. 2a) or time lags (Fig. 2b) are alone
insufficient to explain current ES supply. Rather,
historical land-use produces a ‘‘legacy lock-in’’ that
commits the landscape to consequences for ES that do
not readily respond to contemporary intervention. We
illustrate several examples where threshold dynamics
can cause rapid and surprisingly persistent change in
ecosystem services in US cultural landscapes.
Hydrologic ES (Brauman et al. 2007) in cultural
landscapes may be especially vulnerable to threshold
dynamics associated with past land use and resistant to
reversal. In agricultural landscapes, long-term use of
nitrogen (N) fertilizers is a ubiquitous cause of
groundwater contamination (Di and Cameron 2002),
as discussed in the previous section. Groundwater
nitrate concentrations may increase gradually over
time, and rural wells can exceed threshold concentra-
tions ([3 mg/l) known to result from anthropogenic
activities and to be dangerous to human health, e.g.,
associated with blue baby syndrome (US EPA 1996;
LaMotte and Greene 2007; Tesoriero and Voss 1997).
Nitrate is the most widespread groundwater contam-
inant in Wisconsin, and groundwater nitrate concen-
trations have increased in rural Wisconsin (Saad
2008). In 2015, 20–30% of freshwater wells in
south-central Wisconsin could not provide water for
human consumption because nitrate concentrations
exceeded the maximum contaminant level (Meche-
nich 2015). There is concern that nitrate concentra-
tions are poised to increase further as nitrate penetrates
into deep aquifers and moves farther from original
source areas (Kraft et al. 2007). Whether elevated
nitrate concentrations produce legacy lock-ins will
depend on the balance between groundwater recharge
and output. However, groundwater turnover times are
typically very slow, making it difficult to reverse water
quality degradation.
In similar vein, P enrichment degrades lake water
quality and myriad ES associated with lakes across
agricultural, urban, and ex-urban cultural landscapes
Landscape Ecol
123
(Carpenter et al. 1998, 2006; Lathrop 2007; Carpenter
and Lathrop 2008) and is associated with slow
processes, as discussed above. This bank of soil P
guarantees high future loadings to lakes, even if
fertilizer application ceases (Carpenter and Lathrop
2008; Kara et al. 2011). However, once a lake
becomes eutrophic, additional threshold dynamics
may entrain the lake in the turbid state, even if P inputs
cease. Specifically, lakes of intermediate depth are
least reversible because internal P recycling from
sediments cannot be mitigated by aquatic macro-
phytes, so the lake remains in the undesirable
eutrophic state (Genkai-Kato and Carpenter 2005).
Thus, land-use legacies associated with slow pro-
cesses can interact with the abiotic template (e.g., lake
morphometry) in ways that make it difficult to
improve the hydrologic services associated with
high-quality surface water once a threshold has been
passed.
Flood mitigation is another hydrologic service that
can show threshold dynamics and lead to legacy lock-
in. In urbanizing landscapes, increase in impervious
surfaces (e.g., roads, parking lots, roofs) associated
with expanding development reduces the capacity of
the landscape to absorb rainfall, which then exacer-
bates flooding. Short-term increases in runoff volume
and rate following rain events (i.e., flashiness; Poff
1996) rise with extent of impervious surface, and early
studies suggested a threshold of *15% impervious
surface was associated with undesirable ecosystem
responses including flooding (Paul and Meyer 2001).
Impervious surface cover increased between 1916 and
2013 in the watersheds of two urban lakes in
Wisconsin, reaching 12.6 and 28.9% for Lakes
Mendota and Monona, respectively; flashiness also
increased, and impervious surface was the strongest
driver (Usinowicz et al. 2017). Greater flashiness
indicates more potential flooding, which is increas-
ingly observed in these watersheds. If the frequency of
high-intensity rainfall increases, even areas of the
landscape that did not experience flooding historically
may flood in the future (Usinowicz et al. 2017). The
negative effect of land-use legacies on flood mitiga-
tion may be undetected until the next intense rain
event occurs, revealing an ecosystem services debt.
Climate change is expected to exacerbate such effects
by increasing the frequency and intensity of rain
events in Wisconsin (Wisconsin Initiative on Climate
Change Impacts 2011), and these land-use legacies
may becomemore pronounced in the future. Historical
development patterns may lock-in vulnerability to
flooding for a long time because it is difficult to reverse
patterns of urban and suburban development. Declines
in flood mitigation are theoretically reversible by
replacing impervious surfaces with semi-natural land
cover, but cities are seldom deconstructed.
The degree to which other kinds of ES may respond
to thresholds associated with land-use legacies is not
well known (Burgi et al. 2017). Some threshold
responses, such as those related to size or connectivity
of semi-natural vegetation in cultural landscapes, may
be more amenable to reversal. For example, croplands
could be strategically converted to semi-natural veg-
etation that would support wild pollinators or natural
enemies more readily than cities can be deconstructed.
Of particular importance, however, is the potential for
changing environmental drivers to dampen or amplify
effects of land-use history on ecosystem services. As
environmental change progresses, thresholds that
were not apparent previously may be passed as drivers
change—as more land is developed, temperatures
warm, or hydrologic variability increases. The inter-
action of land-use legacies with changing drivers has
the potential to alter ecosystem services in cultural
landscapes in surprising ways, often over timeframes
beyond those seen in the previous conditions.
Discussion
Persistent effects of past land use on ES supply
permeate cultural landscapes in the US andmany other
regions of the world. We have suggested three
interconnected conditions whereby land-use legacies
are important for interpreting current ES supply, and
illustrated these with examples from agricultural,
urban, and exurban cultural landscapes in the US.
These settings reflect the dominant land-use changes
over the past 100–150 years. Identifying mechanisms
and consequences of these legacies requires attention
to temporal dynamics, including rates of change in ES
and time elapsed since land-use change. In some cases,
ES dynamics may be driven by system variables that
change slowly, leading to lagged responses of ES to
land-use change. In other cases, ES may respond
rapidly to land-use change but effects persist for long
time periods, or changes in ES may exceed a threshold
Landscape Ecol
123
such that recovery to a prior condition is no longer
possible.
The three ES/land-use legacy conditions we pre-
sented here are also amenable to exploration and
testing in regions outside the US. For example, land-
use legacies have been shown to influence above-
ground carbon storage, recreational services, and bird
diversity in urban Sheffield, UK (Dallimer et al. 2015);
examples of extinction debts are prevalent throughout
European landscapes (Dullinger et al. 2013); and
historical grassland management influenced contem-
porary plant biodiversity in a rural Swedish landscape
(Cousins and Eriksson 2002). Application of our
framework in other regions is an interesting direction
for future study, and may yield new insight into the
mechanisms by which historical land-use shapes ES
supply. For example, many studies we have high-
lighted here report legacies from past agricultural use.
Studies documenting legacies from alternate land uses
(e.g., forest harvest or clearance) are comparatively
rare in the cultural landscapes we consider.Whether or
not this trend is generalizable outside of the US
remains a question amenable to further testing.
What are the implications of land-use legacies for
future ES supply? Our examples highlight the influ-
ence of past land use on contemporary ES supply, but
these examples also suggest that today’s land-use
decisions will influence future ecosystem services. For
example, current agricultural practices continue to
enrich soils with P and N. These nutrients will
influence water quality and associated ES in tomor-
row’s cultural landscapes, particularly in the absence
of new strategies to mitigate erosion (Heathcote et al.
2013). Similarly, ongoing habitat loss and fragmenta-
tion will continue to exert lagged effects on biodiver-
sity-based ES, and current expansion of impervious
surfaces risks ‘‘locking in’’ future cultural landscapes
to more frequent flooding. As researchers and policy-
makers increasingly explore possible futures using
scenario analyses (Peterson et al. 2003; Polasky et al.
2011), missing the potential for current land-use
decisions to exert legacy effects will compromise
our ability to project future ES supply. Thus, it is
imperative that rates of processes, potential for time
lagged effects, and thresholds and reversibility be
considered today to manage landscapes sustainably in
the decades ahead.
As we consider future legacies in cultural land-
scapes, wherein lie key uncertainties and research
needs? We highlight three areas we expect to be of
increasing importance in US cultural landscapes in the
years ahead. First, urban landscapes may themselves
generate future land-use legacies that will become
increasingly prevalent as cites change. In this study,
we emphasized effects on ES of transitions from
natural/semi-natural landscapes or agricultural use to
urban or exurban lands. However, as urban popula-
tions expand in many places and contract in others,
legacies of urban land abandonment will become
increasingly common (Nassauer and Raskin 2014).
Examples are already emerging in US cities with high
urban vacancies (Dewar and Thomas 2012), such as
‘‘rust belt’’ cities like Detroit, Michigan that have been
changed fundamentally by global shifts in manufac-
turing. In 2010, Detroit had shrunk to a population of
\714,000 from a 1950 peak of 1.8 million (US Census
Bureau 2010). The resultant urban vacancies occupy a
combined area larger than all parks and open spaces in
the city—an area roughly the size of Manhattan, New
York (Detroit Future City 2013). Such urban shrink-
age provides an opportunity to increase ES provision
in and around urban cultural landscapes (Haase et al.
2014). However, sustainable management of ES on
abandoned land requires understanding the legacies
left behind. Abandoned urban land does not easily
‘‘return to nature’’, as is often construed by urban
residents and the popular press (e.g. characterization
of abandoned lots as ‘‘urban prairie’’ or ‘‘na-
turescapes’’; Ager 2015); altered site hydrology and
soil characteristics remain, even following proper
demolition of urban structures (Nassauer and Raskin
2014). In the US, such legacies are often compounded
by the prevalence of illegal dumping (e.g., of house-
hold chemicals, construction debris, and oil and gas
products) on vacant land (Beauregard 2012). As urban
abandonment and renewal continue, uncertainty sur-
rounding legacy contamination will be a common
challenge for management of ecosystem services in
cultural landscapes.
Second, the long-term consequences of new con-
taminants may present unanticipated challenges for
future ES supply, creating legacies not yet apparent in
today’s cultural landscapes. Microplastics, for exam-
ple, have recently emerged as a threat to marine and
freshwater ecosystems (Cole et al. 2011; Eerkes-
Medrano et al. 2015). Yet, how, and when, microplas-
tics may affect aquatic ES is not well understood.
Similarly, while microplastics are expected to alter
Landscape Ecol
123
physical soil properties and increase concentrations of
soil contaminants (Rillig 2012), their impact on
terrestrial ES is also unknown. What other materials
or practices are being introduced in cultural land-
scapes today that might have consequences that
manifest much later in time?
Third, consequences of land-use legacies on ES
supply may change qualitatively as broad-scale
drivers continue to change. In particular, shifting
climatic drivers may become increasingly important in
modulating effects of historical land use on future ES.
For example, higher intensity and more frequent storm
events predicted due to climate change may mobilize
sediment to a greater extent than in the past, exacer-
bating effects of historical land use on water quality in
cultural landscapes (Carpenter et al. 2015a). Climate-
driven sea-level rise may also exacerbate legacy
effects on ES, most notably in low-lying coastal
regions that are home to a significant portion of the US
population (NOAA 2014). Combined with a projected
increase in storms like Hurricane Sandy, which
flooded major sections of the New York Metropolitan
area in 2012, sea-level rise is likely to worsen flooding
associated with future storm surges in New York (Lin
et al. 2016). Some coastal urban areas, such as
Broward County in southern Florida, are already
considering how to proactively adapt urban infras-
tructure to reduce future flooding (Broward County
Climate Change Action Plan 2015). A long history of
industrial land use in many coastal regions also means
that sea level rise could mobilize legacy contaminants
and threaten water quality and human well being
(Duke et al. 2014). Warming sea surface temperatures
also may increase mercury concentrations in fish,
subsequently increasing human exposure when sea-
food is consumed (Dijkstra et al. 2013). Thus, future
research and landscape planning must consider poten-
tial influences of land-use legacies not in isolation, but
in conjunction with additional drivers of ES supply.
We have presented three general conditions for
when land-use legacies are likely to affect ES supply,
yet we recognize characteristics of land-use history
not covered here will also be important. For example,
the intensity of historical land use, its duration, and the
sequence of past land-use transitions can all amplify or
dampen the magnitude of effects on ES supply (i.e.
modifying the strength of the responses in Fig. 2), but
these characteristics were not explicitly addressed in
our synthesis. Data availability and quality will
influence the ability to detect relationships between
historical land use and ES supply, and the scales of
historical data and ES estimates must align. Addition-
ally, our examples of particular ES are illustrative, and
there are many ES for which the relative role of
historical vs. contemporary landscapes is unknown.
Lastly, few studies consider effects of land-use
legacies on multiple services, or ES bundles (but see
Renard et al. 2015; Tomscha and Gergel 2016). With
the ES field moving towards understanding interac-
tions between services and management implications
(Bennett 2017), future studies should also consider the
role of historical land use in shaping ES relationships.
As ES research becomes increasingly mechanistic,
further consideration of temporal dynamics and land-
use legacies in cultural landscapes is sorely needed. If
a function or process underpinning an ES is affected
by land-use legacies, the service will be affected as
well. Thus, understanding historical land use can be
critical for accurately explaining variation in contem-
porary ES in cultural landscapes and anticipating
future ES supply. Improved understanding of the role
of history will not only lead to better maps and
management of ES supply, but will also aid in decision
making as new challenges to sustaining our cultural
landscapes arise.
Acknowledgements We thank Matthias Burgi for inviting us
to develop this paper, and four anonymous reviewers for
constructive feedback on an earlier version. We acknowledge
funding from the US National Science Foundation, especially
the Long-term Ecological Research (DEB-1440297 and DEB-
1440485) and Water, Sustainability and Climate (DEB-
1038759) Programs, and support to MGT from the University
of Wisconsin-Madison Vilas Trust. CZ acknowledges support
from a Natural Science and Engineering Research Council of
Canada doctoral fellowship.
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